Plastic sex-roles in the common goby – the effect of nest availability

One of the most important factors shaping animal sex-roles is the operational sex ratio, since a skew of this ratio promotes mating competition in the more abundant sex. In this study, we wanted to see if a change in sex-roles, related to an environmental induced change in operational sex ratio, could be demonstrated in a field situation. Common goby ( Pomatoschistus microps ) males build nests under bivalve shells and provide exclusive parental care. Therefore, available nest sites are crucial for successful breeding in this species and should also affect the operational sex ratio. We addressed the question of whether the availability of nest sites, through an effect on the operational sex ratio, affects courtship and mating competition in the common goby. We increased nest site availability in the field by adding potential nest material (bivalve shells) to one location. We then compared the reproductive behaviour of the gobies at this and another location with lower nest site abundance. In general, the reproductive behaviour differed as predicted. Under nest shortage, males occupied new nests and received eggs faster than under nest excess. Behavioural observations of nests revealed that males initiated courtship more often than females where nest sites were abundant, whereas female courtship dominated where nests were scarce. Males more often rejected females attempting to enter their nests under nest shortage, male sneaking attempts were also more common in this bay. Male-male aggression was frequent at both locations, whereas agonistic interactions between females only occurred in the nest shortage bay. Thus, males should be subject to sexual selection in both bays even though courtship roles differed. These results suggest that nest site availability affects mating competition and courtship roles in the common goby.     

Mate choice and cannibalism in a natural population of a stream goby,Rhinogobius sp.

Breeding ecology of the stream goby,Rhinogobius sp. LD (Large Dark), was investigated under natural conditions. Males selectively courted females of similar size to lead them to the nests, whereas females followed courting males preferentially when the relative male size was greater. Male-male competition for a female was relatively infrequent and not severe. Developmental stages of eggs and egg numbers in one nest indicated that males receive 1–3 clutches during one breeding cycle. Males guarding multiple clutches frequently ate some of the eggs, but those guarding single clutches rarely did so. Gravid females in the nest also frequently cannibalized eggs laid by a previous female, probably in order to extend the area available for egg deposition. Mate choice in this species is discussed in relation to paternal ability, limitation of available spawning area and the female-biased sex ratio.     

Intruders in Nests of the Spotless Starling: Prospecting for Public Information or for Immediate Nesting Resources?

The prospective behaviour for nests by conspecific intruders may be a strategy to gather public information for future reproduction or to secure resources for immediate reproduction. Although the relationship between the sex, age and breeding experience of intruders and the sex and age of residents may be crucial for understanding the significance of nest‐prospecting behaviour, a precise determination of these traits has rarely been addressed in field studies. In a free‐ranging population of spotless starlings ( Sturnus unicolor), inexperienced birds were the predominant intruders. The ratio of male‐to‐female intruders was higher among birds without previous breeding experience, although our results did not allow us to determine whether more inexperienced males or females intruded nests. The average annual rate of intrusion of either sex was strongly correlated with the number of fledglings produced in the study colony the previous year, which seems to indicate that the inexperienced intruders were, in general, yearlings. The proportion of the sexes among the intruders in each study year was correlated with the average age of male owners but not with the age of female owners. This result suggests that the presence of intruding males in the nests was affected by an increased opportunity to find vacancies in nesting territories as resident males became older. This finding supports the hypothesis that, in most cases, birds visited nests to obtain personal information about nesting resources. However, the intrusion rates were higher when there were nestlings than during the incubation period, which suggests that collecting public information was also involved in the intruding behaviour. Both sexes regularly attacked intruders, which implies that the intruders inflicted some kind of cost on the owners. Female floaters were more frequently chased, probably because they regularly lay parasitically in conspecific nests.     

Re-use of courtship nests for quick remating in the polygynous Fan-tailed Warbler Cisticola juncidis

In a population of Fan-tailed Warblers Cisticola juncidis near Osaka, Japan, the males were polygynous, building a succession of elaborate courtship nests within their territories, and advertising them to females. Nest desertion by females frequently occurred early in the breeding cycle. Deserted nests were generally left untouched, but 17 out of 88 territorial males cleaned and re-used 20 of 79 nests deserted by females for their next courtship. The frequency of nest re-use increased with season. Of the re-used nests six (300%) were re-occupied by secondary females. Newly built nests were preferred to the 'second-hand' nests, probably because new nests were likely to last longer. Breeding success did not differ between the two categories of nests. In males, nest re-use seems to save time and energy expenditure for additional mating.     

Sex ratio of North Island kaka (

The sex ratio of the kaka population inhabiting the Waihaha Ecological Area, Pureora Forest Park was estimated between late October 1994 and January 1995. The observed sex ratio estimate was three males to one female compared to a capture rate (using mist nets) of six to one in the same area between January and June 1994. Females appeared to be less susceptible to capture than males. The skewed sex ratio toward male kaka was significant and suggests that female kaka suffer higher mortality (probably due to predation at the nest) than males. The slow breeding rate of kaka combined with a high loss of breeding females is a serious threat to the long term survival of the kaka population within the Waihaha Ecological Area.     

Reconnaissance for future breeding sites by spotted sandpipers

We studied the sex-role-reversed, polyandrous spotted sandpiper(Actitis macularia) from 1974 to 1990 in northern Minnesota,USA. After peak arrival of breeding birds and before peak departureat the end of the breeding season, there were many short-termvisitors (transients) to the study site. Stepwise discriminantfunction analysis (DFA) was used to determine the importanceof absolute sex ratio (males/female), sex of the transient bird,number of nests, and number of breeding males and females duringthe week of visit in predicting whether a visiting bird wouldreturn the following year. In addition, multiple regressionwas used to determine how much variability in the number oftransient birds returning in subsequent yearscould be explainedby annual values during the year of transience for numbers ofbreeding males and females, numbers of eggs laid and hatched,and absolute sex ratio. Annual recruitment of foreign adultsranged from 1 to 20 birds, of which 0–56% were seen visitingin previous years. Female recruits were more likely than malesto have been observed previously as transients. Twenty-two chickshatched at our study site returned and bred for the first timemore than 1 year after hatching. Of these, 9 (41%) were seenas transients between the year of hatch and breeding. The DFAshowed that transient females returned more often than transientmales and that the number of transients returning in subsequentyears was positively associated with the absolute sex ratioduring the week visited. When the sexes were analyzed separately,none of the weekly variables significantly discriminated femalereturn, but sex ratio was positively associated with male return.Regression showed that the number of transient birds returningin subsequent years was positively associated with the numberof male breeders at our study site during the year a bird visited.Percentage return the year after transience was positively associatedwith the number of eggs laid at our study site during the yeara bird visited. When sexes were analyzed separately, the higherthe number of female breeders during the year a bird visited,the greater the number of males returning in subsequent years,and greater numbers of breeding males were positively associatedwith transient female return. Based on our results, we suggestthat transient birds were searching for better breeding areasfor future breeding and that intrasexual competition made thisinformation more important to females than to males.     

Successive nest building and polygyny of Fan-tailed Warblers Cisticola juncidis

The polygynous mating system of the Fan-tailed Warbler Cisticola juqcidis was investigated between 1978 and 1981. The male warbler builds many nests unaided; however, he has no more than one active vacant nest for courting at any time. Nest building lasted from April until August. After completing the outer fabric of a nest, the male advertised it and led a female to the nest site by a unique invitation flight. On average a male built 6.5 nests during one breeding season and three of them were accepted by females. The most successful male completed 18 nests, and mated with 11 females. Out of a total of 111 males which established a territory, 30 had no mate, 14 were monogamous, and the rest were polygynous. About 50 to 70% males were polygynous over the four years. The sex ratio varied from 1.41:1 to 2.17:1 (females: male) in the breeding population. It was partly caused by the presence of 'floating males'. After the completion of the outer fabric of the nest, the male warbler did not take any further role in nesting and caring for young.
The polygynous mating system of the Fan-tailed Warbler is characterized by successive nest building. Its extreme development results from the long breeding period and the male having no role in parental care.     

Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor

The sexes’ share in parental care and the social mating system in a marked population of the single‐brooded Lesser Spotted Woodpecker Dendrocopos minor were studied in 17 woodpecker territories in southern Sweden during 10 years. The birds showed a very strong mate fidelity between years; the divorce rate was 3.4%. In monogamous pairs, the male provided more parental care than the female. The male did most of the nest building and all incubation and brooding at night. Daytime incubation and brooding were shared equally by the sexes, and biparental care at these early breeding stages is probably necessary for successful breeding. In 42% of the nests, however, though still alive the female deserted the brood the last week of the nestling period, whereas the male invariably fed until fledging and fully compensated for the absent female. Post‐fledging care could not be quantified, but was likely shared by both parents. Females who ceased feeding at the late nestling stage resumed care after fledging. We argue that the high premium on breeding with the same mate for consecutive years and the overall lower survival of females have shaped this male‐biased organisation of parental care. In the six years with best data, most social matings were monogamous, but 8.5% of the females (N=59) exhibited simultaneous multi‐nest (classical) polyandry and 2.9% of the males (N=68) exhibited multi‐nest polygyny. Polyandrous females raised 39% more young than monogamous pairs. These females invested equal amounts of parental care at all their nests, but their investment at each nest was lower than that of monogamous females. The polyandrously mated males fully compensated for this lower female investment. Polygynous males invested mainly in their primary nest and appeared to be less successful than polyandrous females. Polyandry and polygyny occurred only when the population sex ratio was biased, and due to strong intra‐sexual competition this is likely a prerequisite for polygamous mating in Lesser Spotted Woodpeckers.     

The effect of nest aggregation on the reproductive behaviour of the peacock blenny Salaria pavo

The effect of nest aggregation in courtship behaviour was tested experimentally in an ecologically constrained, sex-role reversed population of the peacock blenny Salaria pavo . Mixed sex groups of eight males and eight females were tested in experimental tanks, containing eight potential nests either aggregated or dispersed. In the aggregated treatment, males spent more time inside their nests and monopolized other potential nests, causing a female-biased operational sex ratio (OSR). In the aggregated treatment, females also expressed more courtship behaviour. The results in general support the prediction that the aggregation of nests promotes male monopolization of potential nests, resulting in fewer nest-holding males and therefore a female-biased OSR that leads to the reversal of sex roles.     

Brood desertion and polygamous breeding in the Kentish Plover Charadrius alexandrinus

In the Kentish Plover Charadrius alexandrinus one of the adults, typically the female, deserts the brood when the chicks are a few days old. Once parental care is terminated, adults may initiate a second nesting attempt if sufficient time remains within the season. For these nests, individuals pair with different mates from those of the first nesting attempt, thus becoming sequentially polygamous. In a small population of Kentish Plovers in Fuente de Piedra lake (southern Spain), the duration of biparental care of broods was longer than in other localities. It also showed considerable variation between years that was evidently related to Gull-billed Tern Gelochelidon nilotica predation pressure on the chicks. There was year-to-year variation in the number of polygamous matings. Both the duration of the breeding season and nesting success in the first half of the season limited the occurrence of polygamy. Despite females deserting broods earlier than males, the interval between the first and second nesting of polyandrous females and polygynous males was similar. The interval was not affected by the body condition of females after the first nesting attempt, nor by problems related to egg formation ability, but was probably due to the availability of potential mates. More females than males initiated second nests, suggesting that polygamous opportunities were more limited for males than for females. In terms of delayed breeding, reduced survivorship or reduced breeding opportunities in years following polygamous breeding, polygamous individuals did not have greater costs than non-polygamous ones. Females with second nests did not seem to be selective in mate choice, mating with any available male. Mates for second nests may therefore be of lower quality than those for first nests, as judged by male plumage characteristics. Clutch sizes and egg characteristics of polyandrous females were similar in first and second nests. Nest success of second nests was only 40% of that of first ones, with nest desertion accounting for 60% of the losses. As the costs of polygamy are apparently low and as breeding success is very variable among years, polygamous breeding of the long-lived Kentish Plover may be an important breeding strategy with which to increase individual lifetime reproductive success.     

Age dependent sex allocation in the polygynous spotless starling

Theoretical and empirical evidence suggests that avian females are able to manipulate the offspring sex ratio at birth. Although mating with an attractive male may induce females to skew the sex ratio toward males, the balance between the benefits of producing attractive sons and the costs of competing with other females for mates could vary with female age, a possibility that had not been previously explored. In this paper we increment experimentally the attractiveness of males of the polygynous spotless starling (Sturnus unicolor) by adding green plants to their nests, a trait involved in courtship, and look for female age-differential effects on offspring primary sex ratio. Young and middle aged females produced more sons in experimental than in control nests, as expected, but old females showed the opposite tendency. To explain this novel result, we speculate that older females are limited to produce the most costly sex because the physiological drawbacks imposed by ageing reduce their ability to compete with younger ones for the non-shareable resources offered by males. We discuss that this evolutionary scenario may be widespread in avian polygynous systems.     

Effects of sex ratio on male behaviour and reproductive success in a field population of threespine sticklebacks (Gasterosteus aculeatus) (Pisces: Gasterosteidae)

Biased sex ratios of breeding threespine sticklebacks (Gasterosteus aculeatus L.) occur naturally in tide-pools of a Quibec saltmarsh. We experimentally manipulated sex ratios in certain pools to evaluate the effects on male behaviour and reproductive success (RS). Sticklebacks were stocked at male: female ratios of 1: I, 1: 2 and 2: 1 and observed for a 23-day inter-tidal period. In male-biased pools, only half of the males built nests, compared to nearly 100% in unbiased and female-biased pools. Males in male-biased pools also were less likely to rebuild after losing a nest, visited their nests less often, were more likely to abandon or destroy their nests, had lower RS (measured as the proportion of males hatching fry), but hatched fry sooner, than males in other pools. In female-biased pools, males built nests sooner, lost more nests due to nest-raiding by females, spent more time in aggression (proportion of time spent fighting and threatening), spent more time attacking female conspecifics than male conspecifics or heterospecifics, and courted more frequently, than males in other pools. Habituation to conspecific males, but not to females, occurred in all pools. These findings are discussed with respect to sexual selection theory.     

Few, small, and male: multiple effects of reduced nest space on the offspring of the solitary wasp, Euodynerus (Pareuodynerus) posticus (Hymenoptera: Vespidae)

The size of the preexisting wood cavities used as nests by aculeate Hymenoptera is expected to have consequences on fitness parameters such as offspring number and size. We evaluated the consequences of using small and large (three-times more voluminous) trap-nests by the solitary wasp, Euodynerus (Pareuodynerus) posticus (Herrich-Schaeffer). Following life-history and sex allocation theories, a number of non-mutually exclusive hypotheses were formulated: i.e. small nests either produce smaller or fewer offspring and/or more males, the cheaper sex. Wasps built about 28% more, but shorter brood cells in large nests, although their volume was still much higher in large nests. Adult males had smaller body size in small nests, but female size did not differ between large and small nests, possibly as an adaptive response against the future higher foraging costs of size-reduced females. Sex-ratio was often biased towards males in small nests. Mortality did not differ between large and small nests. We conclude that E.?(P.) posticus females would benefit from using larger nests, but that the sex-ratio would be probably overall unbalanced if females would not use also smaller, male-oriented tunnels.     

Cross‐fostering eggs reveals that female collared flycatchers adjust clutch sex ratios according to parental ability to invest in offspring

Across animal taxa, reproductive success is generally more variable and more strongly dependent upon body condition for males than for females; in such cases, parents able to produce offspring in above‐average condition are predicted to produce sons, whereas parents unable to produce offspring in good condition should produce daughters. We tested this hypothesis in the collared flycatcher (Ficedula albicollis) by cross‐fostering eggs among nests and using the condition of foster young that parents raised to fledging as a functional measure of their ability to produce fit offspring. As predicted, females raising heavier‐than‐average foster fledglings with their social mate initially produced male‐biased primary sex ratios, whereas those raising lighter‐than‐average foster fledglings produced female‐biased primary sex ratios. Females also produced male‐biased clutches when mated to males with large secondary sexual characters (wing patches), and tended to produce male‐biased clutches earlier within breeding seasons relative to females breeding later. However, females did not adjust the sex of individuals within their clutches; sex was distributed randomly with respect to egg size, laying order and paternity. Future research investigating the proximate mechanisms linking ecological contexts and the quality of offspring parents are able to produce with primary sex‐ratio variation could provide fundamental insight into the evolution of context‐dependent sex‐ratio adjustment.     

Seasonal sex role changes in the blenniid Petroscirtes breviceps, a nest brooder with paternal care

The plasticity of the sex roles in the blenniid fish Petroscirtes breviceps , a nest brooder with exclusive paternal care, was studied throughout an 8 month breeding season. Males performed most courtships early and late in the breeding season, whereas females performed most in the middle of the season. These results indicated that the sex of individuals initiating courtship changed seasonally, with courtship role reversal in the middle of the season. Intrasexual aggression in both sexes occurred much more frequently in mid-season than in the early and late seasons. Males frequently fought when available nest sites were limited, regardless of the presence of females, suggesting that males competed for nests in order to qualify to mate (resource competition). In contrast, courting females fought only in mid-season, when females' relative success in entering nests decreased, indicating that females competed for limited mating opportunities (mating competition). The reversed courtship roles and female mating competition in mid-season suggested that the sex roles in P. breviceps changed seasonally from the conventional roles to reversed roles and back again during one breeding season. This study provides the first empirical evidence of multiple changes in the sex roles of animals within a breeding season.     

Factors limiting polygyny in the willow ptarmigan

Willow ptarmigan (Lagopus lagopus) usually pair monogamously and males invest more in vigilance and parental care than do males of other grouse species. I tested whether polygyny is rare because male parental care is necessary for successful reproduction. By continuous removal of males I skewed the operational sex ratio, induced polygyny, and then compared breeding success and survival of females that shared a mate and those that did not. Both groups had similar clutch sizes, laid eggs at the same time, produced young of the same weight, and fledged similar numbers of juveniles. Hens that shared a mate suffered higher losses of nests during incubation, and fewer returned to the breeding range in subsequent years. Thus, although male parental care was not essential, it did improve female reproductive success and survival. I suggest that willow ptarmigan are not polygynous because territorial females are able to prevent potential secondary hens from settling, thus securing unshared access to male investment and territorial resources.     

Intranidal mating,emergence, and sex ratio in a communal beeAndrena jacobi Perkins 1921 (Hymenoptera: Andrenidae)

The extent of preemergence intranidal mating, schedules of emergence, and sex ratio at emergence were documented forAndrena jacobi, a communal, univoltine bee, by collecting and dissecting adults as they emerged from their fossorial nests in 1994. Over 70% of females had mated intranidally with nestmate males, thus potentially incestuously, before emerging. Preemergence intranidal mating did not vary during a day or between nests within a day, though it was less frequent at the start and end of the period of emergence. It was independent of a female's size.A. jacobi was protandrous, though some males emerged after all females. The sex ratio at emergence was remarkably female biased, possibly a consequence of local mate competition. Intranidal mating may represent a characteristic trait of communal bees where a high density of receptive females are predictably aggregated within a nest.     

Factors affecting female-biased sex ratio in a trap-nesting wasp, Trypoxylon malaisei

We examined the female-biased sex ratio of a trap-nesting wasp Trypoxylon malaisei considering the following factors: (1) local mate competition (LMC), (2) resource quality, (3) partial bivoltinism, and (4) presence of constrained females. The sex ratio (expressed as male ratio) at emergence was strongly female biased, i.e., 0.30 and 0.19, in terms of the number and investment, respectively. To evaluate the primary sex ratio, we analyzed the data from nests where all the offspring successfully emerged, excluding nests composed of single-sex offspring. The primary sex ratio was also female biased, at 0.33 and 0.21, in terms of the number and investment, respectively. LMC was highly responsible for the female-biased sex ratio because both the nonrandom oviposition sequence [females at inner cells and male(s) at outer cells] and earlier emergence of males allowed sib-matings to occur. In contrast, the other three factors little affected the female-biased sex ratio: the sex ratio was fairly constant when resource quality (nest size) varied, partial bivoltinism was extremely rare or absent, and constrained females were absent or did not reproduce at all. Received: June 19, 1998 / Accepted: January 18, 1999     

Egg sex ratio and paternal traits: using within-individual comparisons

Empirical studies of sex ratios in birds have been limited dueto difficulties in determining offspring sex. Since molecularsexing techniques removed this constraint, the last 5 yearshas seen a great increase in studies of clutch sex ratio manipulationby female birds. Typically these studies investigate variationin clutch sex ratios across individuals in relation to environmentalcharacteristics or parental traits, and often they find no relationships. In this study we also found that clutch sex ratiosdid not vary in relation to a number of biological and environmentalfactors for 238 great tit Parus major nests. However, interestingsex ratio biases were revealed when variation in clutch sexratios was analyzed within individual females breeding in successiveyears. There was a significant positive relationship betweenthe change in sex ratio of a female's clutch from one yearto the next and the relative body condition of her partner.Females mating with males of higher body condition in yearx + 1 produced relatively male-biased sex ratios, and the oppositewas true for females mated with lower condition males. Within-individualanalysis also allowed investigations of sex ratio in relationto partner change. There was no change in sex ratios of femalespairing with the same male; however, females pairing with anew male produced clutches significantly more female biased. Comparisons of clutch sex ratios within individuals may be apowerful method for detecting sex ratio variation, and perhapsfemale birds may indeed manipulate egg sex but require personalcontextual experience for such decisions.     

Correlates of reproductive success in stream-dwelling male rock bass,Ambloplites rupestris (Centrarchidae)

Synopsis Reproductive success was determined for individual male rock bass, Ambloplites rupestris, inhabiting the Middle Thames River near London, Ontario. Successful males produced numbers of young similar to rock bass breeding elsewhere, but the probability of males producing any young at all was comparatively small. Flooding, predation, and fouling of nests by algae were the major causes of brood mortality. Brood vulnerability was greatest within the egg, when nests were probably most conspicuous to predators. Male reproductive success was positively correlated with large size, early nesting, warm water conditions, and minimal flow rates. Larger males were chosen preferentially by females, and renested more often.